Dwarf mistletoes (Arceuthobium species) are a group of vascular plants that
parasitize conifers. These plants are integral part of forested ecosystems, and have
existed as part of the coniferous forests of North America since the Miocene. Dwarf
mistletoe is important to the ecology of these systems. The fruit, foliage and pollen of
dwarf mistletoe are a food source for numerous bird, mammalian and insect species. Dwarf
mistletoe alters the growth patterns of infected trees, creating structural complexity
within forests in the form of witches brooms and snags, both which are used by numerous
wildlife species for nesting, roosting and cover. These mistletoes are considered serious
pests (by silviculturalists) throughout much of Western North America because they infect,
and can reduce the growth rates of commercially important conifers. In the Southwestern
United States, ponderosa pine is the primary commercially important species infected by
dwarf mistletoe. Land use activities in Southwest forests during the past 125 years have
encouraged the spread of dwarf mistletoes. Many of the silvicultural challenges created by
these parasites are exacerbated by ecologically insensitive land management policies such
as fire suppression, livestock grazing, and inappropriate silvicultural techniques. In
general, dwarf mistletoe only becomes a problem when land managers attempt to create
highly productive forests or tree farms to grow timber far in excess of historical
production rates. The damaging effects of mistletoe can best be minimized, and their
ecological benefits maximized, by recreating forest stands with age, size and density
distributions similar to the original, presettlement forests.

Introduction

Dwarf mistletoes (Arceuthobium species) are an integral component of most
coniferous forests in western North America. Dwarf mistletoe range from central Canada and
southeast Alaska to southern Mexico (Fig. 1). The western United States and northwestern
Mexico are the center of diversity for this genus, containing 24 of the known 32 species
in the world 31. Arceuthobium utilize a wide variety of conifers as their hosts,
although most species prefer pines 31. Two important Arceuthobium species quite
prevalent in the Southwest are A. vaginatum and A. douglasii, which infect
ponderosa pine (Pinus ponderosa) and Douglas fir (Pseudotsuga menziesii),
respectively. This paper examines the ecology of these and other parasite-host complexes,
but is primarily focused on applying this information to the management of Southwest
ponderosa pine forests.

Arceuthobium are well known to most silviculturalists in the Southwest because
they diminish the cash value of Douglas fir and ponderosa pine, the regions most
commercially important conifers60. Accordingly, concerted efforts have been made to rid
forests of these plants on federal lands. However, such a strategy is not wise (or
feasible). Fossil records clearly show that Arceuthobium species have been
associated with North American coniferous forests since the Miocene (12-26 million years
BP) 39. Over this vast expanse of time, many species have adapted to and are perhaps even
dependent on mistletoes. Numerous species of birds, mammals and arthropods have been
observed feeding on the fruits and foliage of mistletoe, while other birds and mammals
utilize the witches brooms created by the mistletoe as nesting platforms or roosting sites
(Appendices 1 and 2). Witches brooms are abnormal branches found on trees infected by Arceuthobium.
These brooms contain dense patterns of interconnecting twigs and branches, forming a
unique structural element in the forest that has many ecological benefits.

Although dwarf mistletoe and witches brooms are integral components of Southwest
forests, the primary focus of forest managers in the National Forests has been to
eliminate them. Because of the agencys overwhelming focus on timber production, most dwarf
mistletoe research sponsored by the Forest Service has been focused on control methods,
with little attention being paid to their ecological importance (e.g. see symposia
proceedings by Scharpf and Parmeter 1978, Hawksworth and Scharpf 1984). Dwarf mistletoe
control projects have traditionally been pursued by the Forest Service in order to
maximize timber production. As a result, attention has been focused on management
techniques to produce more merchantable trees at the expense of managing for ecosystem
health. There is an urgent need for the Forest Service to reevaluate its current strategy
for managing dwarf mistletoe, and to adopt an integrated ecosystem perspective that
manages for forest ecosystem integrity, rather than waging a war (sensu Wicker, 1984)
against dwarf mistletoe 74. In this paper we examine three crucial issues related to dwarf
mistletoe in Southwest forests in the hopes of bringing a more enlightened approach to the
management of these forests: (1) The biology and ecological importance of dwarf mistletoe,
(2) how fire suppression, livestock grazing and silvicultural practices have altered the
interaction of dwarf mistletoe with its environment, and (3) integrated, ecosystem-based
strategies for managing dwarf mistletoe.

The Importance of Dwarf Mistletoe to the Ecology of Southwestern Forests

Dwarf mistletoe interacts with other plants and animals in Southwest forests in several
ways: (1) their fruit and foliage provide a food source to animals (2) the brooms they
help form adds short-term structural complexity to the forest, and (3) by hastening the
death of trees, they help to create snags, which are an important source of long-term
structural complexity and wildlife habitat. Additionally, by influencing the amount of
dead wood in a forest, they indirectly affect fire intensities, and thus the successional
dynamics of forests. This latter topic will be covered under Fire and Dwarf Mistletoe.

Consumers of Dwarf Mistletoe
There are a number of animals that have been observed eating the fruit and foliage of
dwarf mistletoe (Appendix 1). Such species include elk, deer, squirrel, grouse, pygmy
nuthatches and mountain chickadees, as well as a number of arthropods 7-11, 16, 23,
40, 71-74. The relative importance of dwarf mistletoe as a food source for such
species is not known, but given the prevalence of mistletoe, it is likely that certain
species have come to rely upon it as a food source during certain times of the year. It is
known that the foliage of dwarf mistletoe provides a source of food in the winter for
grazers, when fresh foliage is scarce. Witches brooms create ideal platforms for trapping
snow, making them susceptible to breakage. These breakages provide mistletoe foliage to
animals such as elk and deer 9. Nutritional analyses of dwarf mistletoe
indicate that the foliage and berries have high nutritional value for deer, and likely for
other species as well 73. Dwarf mistletoe also helps to provide food for other
species early in the spring. Branches of conifers such as Douglas fir, that are infected
with dwarf mistletoe, break bud (bud out) earlier in the spring than do uninfected twigs,
providing a food source for certain insect groups 70, which in turn provide
food for insectivores.

Dwarf Mistletoe as Food
forWildlife

For numerous birds and mammals, dwarf mistletoe is an
important part of their diet. The foliage and berries of these plants are highly edible.
Twelve wildlife species have been documented in the scientific literature as eating dwarf
mistletoe, including game animals such as elk, deer and grouse. It is known that dwarf
mistletoe is highly nutritious, and if more research efforts were put towards learning
about their ecology (rather than studying ways to eliminate them), we would likely find
that many more wildlife species use these plants as a food source.

Witches Brooms as Wildlife Habitat
The habitat created by witches brooms is important to a number of birds and mammals.
Forests infected with mistletoe have been strongly correlated with the biodiversity of
birds in Ponderosa pine forests 8, in part because of the brooms, which provide
ideal platforms for nesting and cover. It is well documented that birds of prey such as
accipiters and owls use witches brooms as nesting sites 20, 40, 53, 45, 46, 12, 11.
These birds seem to prefer the large brooms found up high in older trees. The formation of
such a broom requires infection of a large tree, because young trees that are infected
tend to die before they reach a large size. These large brooms provide visual protection
from predators both above and below. It is quite difficult to determine whether a broom
contains a nest when viewed from the ground (as many researchers have discovered). A
well-disguised nest in a large broom can also be difficult to detect from the air,
providing protection against predators such as the great horned owl 46.

Available evidence suggests that the size of the broom needed for nesting is a function
of bird size 40, 53, 45, 46, 12, 11. Therefore, it is likely that current
logging practices, which preferentially remove large trees, tend to have a
disproportionately larger impact on big birds of prey. This habitat need may partially
explains why two birds of the Southwest which use brooms as nesting sites, the Mexican
spotted owl and the Northern goshawk are listed as endangered and as sensitive,
respectively, by the U.S. Fish and Wildlife Service and the U.S. Forest Service. Several
recent studies have demonstrated that the spotted owl shows a marked preference for
witches brooms as nesting sites 68, 24, 65. Of all the nests examined in these
studies, 153 out of 359, or 43%, were associated with witches brooms 68, 24, 65.
This is particularly remarkable in light of the fact that large brooms of sufficient size
for nesting are rare in Southwest forests. It is quite possible that given an abundance of
brooms to choose from, an even higher percentage of spotted owls would nest in these
structures.

The list of animals known to utilize brooms as nesting platforms is extensive, but not
exhaustive. There have been no thorough studies to determine which Southwestern birds and
mammals use brooms. They may well be crucial habitat for other endangered species. Other
birds known to use brooms as nesting sites include the Coopers hawk, great gray owl,
long-eared owl and others (see inset and Appendix 2 ).

It is well known that species associated with ponderosa pine forests are declining in
numbers and that bird diversity is decreasing. Bird surveys conducted yearly since 1968 in
logged ponderosa pine forests in New Mexico indicate that 75% of all bird species are
declining 44. Overall, about 25% of all species associated with ponderosa pine
forests are declining 21, 32. How much of this is attributable to the loss of
witches brooms is uncertain, but the growing list of birds that use witches brooms as
nesting sites that are also experiencing population declines, suggests that the loss of
this important habitat feature may at least in part be contributing to the loss of
biodiversity in Southwest forests.

Witches Brooms and Wildlife

The witches brooms created by dwarf mistletoe are important nesting
sites for birds and mammals. Scientists have determined that where these structures are
abundant in Southwest forests, bird diversity is very high and where they are absent,
diversity is low. Brooms create an ideal platform for nesting, especially for large birds
such as raptors. The dense cover provided by brooms helps to conceal nests, making them
hard to detect by predators on the ground and in the air. Eighteen wildlife species have
been documented in the scientific literature as nesting in witches brooms, including such
rarities as the Mexican spotted owl and the northern goshawk.

One of the most important ecological functions of dwarf mistletoe is to increase the
rate of snag formation in Southwestern forests. Mistletoe has always existed as a stand
replacing mechanism that has helped to create ecologically important large snags, while
simultaneously opening up the canopy to allow for the establishment of young trees.
Mistletoe infections eventually result in death for many trees, and the average life span
of infected trees is considerably shorter than uninfected trees 29. In general,
infected large trees live longer than infected small trees, although precise relationships
between tree size or age and post-infection longevity are not known. Snags are an
important resource for many species of cavity nesting birds and mammals 63, 52, 48.
The number of snags and cavity nesting birds in Southwest forests are correlated to the
degree of mistletoe infection 8. Much of the recent decline in cavity nesting birds in
southwest forests has been attributed to the loss of large snags. Bird species diversity
and numbers in the Southwest have been positively correlated with the density of mature
live ponderosa pines and snags 3. Snags are used by 85% of North American birds
37, at least 49 species of mammals, and many reptiles, amphibians and
invertebrates 19. Thirty percent of all North American birds nest in snags 47
and forty bird species nest in ponderosa snags 64. Secondary cavity nesters
alone make up 33% of breeding bird species, and 40% of total breeding bird pairs in
ponderosa pine forests 4. Eighty two percent of secondary cavity nesters breed
exclusively in dead and dying trees 4, while between 60 and 94% of
overwintering ponderosa pine-associated birds require snag roosts 69. In
addition to nesting and roosting sites, snags and broken-tops are used as drumming posts,
song perches, hawking platforms and foraging.

Large snags are preferred by primary and secondary cavity nesters. Seventy five percent
of cavity nests on the Coconino National Forest are in trees > 24 inches dbh 17, while
the mean dbh for trees containing cavity nests on the Apache-Sitgreaves National Forest is
23 inches 63. The availability of suitable nesting cavities is the primary
limiting factor in secondary cavity nester populations sizes 2, 28, 13, 51, 76, 5.
Studies demonstrate that where unlimited nesting and roost sites are available, other
factors such as availability of food, do not affect population sizes 41. Mature
forests are the most favorable to cavity nesters because of their abundance of large dead
and dying trees. The removal of these trees in managed forests dramatically decreases the
number and diversity of secondary cavity nesters. The loss of natural bird diversity in
managed forests has been well documented, and the general decline is largely accounted for
in the disappearance of cavity nesting species 27.

Because large trees infected with dwarf mistletoe have been viewed as infection centers
for nearby smaller trees, the Forest Service has promoted a policy of removing these large
trees 55. Such a management strategy removes the next generation of large
snags, leading to the long-term loss of prime rearing habitat for cavity nesting birds.

Fire and Dwarf Mistletoe
Fire was historically an integral component of forest development in the Southwest, and
helped to control the spread of dwarf mistletoe by: (1) keeping overall tree densities
low, (2) selectively destroying infected trees and stands, and (3) pruning infected limbs
from live trees. Additionally, smoke from fires may have inhibited dwarf mistletoe seed
germination.
Today, the number of trees per acre on some southwestern pine forests are one to two
orders of magnitude higher than during pre-European times 15. The reasons for this
increase are discussed below under the Land Use Practices section. High stand densities
contribute to the spread of dwarf mistletoe by placing more trees within the range of
dwarf mistletoe seed dispersal from infected trees.

Estimates of pre-European tree densities range from 9.3 to over 150 trees per acre in
the ponderosa pine type, depending upon growing conditions and method of measurement 14,
15, 38. Forests tended to be more open at lower elevations, on southern exposures,
and on poor soils. Forest were denser in wetter, cooler areas and on richer soils. While a
complete picture of pre-European forests has yet to be established, it is clear that in
many areas, todays ponderosa pine forest is thicker than in the past. As a result,
mistletoe may spread from tree to tree more easily than in the past.

Additional evidence suggests that some ponderosa pine forests had a clumped
distribution, with small clusters of trees imbedded in a grassland matrix 15.
The areas of grass between tree clusters likely formed barriers to the dispersal of
mistletoe seed, helping to limit infections to isolated patches of trees. It must be
emphasized that the density of presettlement southwest forests was quite variable and that
some of the more productive forests were dense enough to facilitate the spread of
mistletoe by direct dispersal. We simply use some estimates of presettlement tree
densities to show that historically, there were many forests where the rate of spread of
mistletoe infections was likely quite low, and infections were not as widespread as they
are today.

There is also evidence to suggest that infected trees are weeded out by fires 1.
Because witches brooms contain highly resinous, flammable material, dwarf mistletoe
influences the susceptibility of trees to fire. Often, brittle brooms break off of trees,
forming a pile of inflammable material at the base of the tree. These broom piles can
support a fire of sufficient intensity to kill a tree. In heavily infected stands, there
may be enough broom piles and dead trees that the entire stand will conflagrate, killing
all trees in a wide area. Such an event, provided it kills all infected trees, serves to
sterilize the area of dwarf mistletoe, keeping it free from this parasite for decades 1.
If some infected overstory trees survive however, they can serve as infection centers,
raining mistletoe down upon the emerging understory of young trees regenerating after the
burn 56. The historical record suggests that such large, stand destroying fires
were rare in ponderosa pine forests 14, and it is likely that most fires
stimulated by the presence of brooms and other dead wood resulting from mistletoe
infections at most killed only single or small clusters of trees.

It also appears that prior to extensive fire suppression, the heat from frequent ground
fires was sufficient to prune back mistletoe infected branches in the lower crowns of
ponderosa pines 57, 58. This served to limit the extent of mistletoe to the
upper reaches of the forest canopy 57, 58. This pruning approach to dwarf
mistletoe control has been successfully employed by modern day foresters to increase the
longevity of infected stands 62.

There is also evidence to suggest that the smoke from fires was deadly to mistletoe
seeds. In an experimental study, researchers found that exposure of mistletoe seeds (A.
vaginatum, A. americanum and A. cyanocarpum) to wood smoke resulted in a germination
rate of less than 5% (as compared to a normal germination rate of 27%) when the seeds were
exposed to smoke for an hour 78. In general, exposure to smoke inhibited
germination after 10 minutes, and virtually no seeds germinated after long exposure times
(90 minutes). Although the importance of smoke to inhibiting dwarf mistletoe seed
germination under natural conditions is not known, this study suggests that it could be
important, especially in light of the documentation by early European explorers that
ponderosa pine forests were often quite smoky during the summer months, when ground fires
were burning 14.

Figure 2. The probability of being infected by mistltoe
seed decreases exponentially as a function of the distance from the seed source (see text
for sources).

The Biology of Dwarf Mistletoe

The Life Cycle of Dwarf Mistletoe and Short Distance Seed DispersalArceuthobium species have a unique and interesting life cycle that is well adapted
to surviving in coniferous forests. The life cycle is illustrated in Figure 2, and has
been well described by Hawksworth elsewhere 31. The summary presented here is a
synthesis of that information. The adult plant produces berries which contain a single
seed. As the fruit matures, fluid pressure builds up inside, until the berry finally
breaks off the stem. At this moment, the released pressure ejects the seed from the fruit
at tremendous velocities, sometimes exceeding 60 miles per hour. Dwarf mistletoe can eject
their seeds up to 100 feet under optimal wind conditions, but in general most seeds travel
no more than 10-15 horizontal feet from their source, and a tree 40 feet from an infected
tree is considered safe from infection. The average, long-term rate of spread for dwarf
mistletoe by such short-distance seed dispersal is about 1-2 feet per year, in managed
stands. The seed contains a viscous coat, which allows it to stick to any object with
which it comes in contact, such as the needles of a conifer. The seeds remain stuck to the
conifer needles until they absorb moisture (e.g. during a rainstorm), whereupon they slide
down to the base of the needle, adjacent to a twig. Young, needle-bearing twigs are most
susceptible to infection. Once in this position, they begin to grow a hypocotyl, or small
root, and bore into the outer layers of the twig. At this stage, swelling occurs in the
infected area on the tree. About two years after the infection, aerial shoots develop, and
two years after that, the female plant develops mature fruit. Dwarf mistletoe are
dioecious (literally, of two houses) meaning that the male and female flowers are found on
separate plants. Although individual shoots of the plant die after four or five years, the
vegetative part of the mistletoe continues to grow, and old colonies are recognizable by
the dense and misshapen cluster of twigs and branches known as witches brooms 30.

Long distance seed dispersal
Although the seeds of mistletoe are spread to nearby trees by direct contact upon
ejection, animals also serve as dispersal agents, and help to explain why mistletoe plants
have been found far from the nearest potential seed source 49. Birds and small
mammals feed on and adjacent to mistletoe as ripe seeds are being ejected, and some of the
seeds attach to their bodies. As the animals move to other trees, the seeds may detach.
Birds such as mountain chickadees and pygmy nuthatches, which forage on the tips of
branches and in the foliage, where seed germination is most likely to be successful, are
thought to be particularly important agents of dispersal 33. Birds apparently
do not transfer viable dwarf mistletoe seeds through their feces. Most such seeds passing
through the digestive systems of birds will not germinate 77, 33. The success
of seeds dispersed by birds is quite low. One study estimated that in a 150 ha plot of
ponderosa pine, there was only one successful long distance infection every four years 33.
A number of birds and mammals, including Stellars jays, red crossbills, Cassins finches,
red squirrels, flying squirrels and pine martens have been observed carrying mistletoe
seeds on their fur or feathers. A complete list of dwarf mistletoe seed vectors is given
in Appendix 3.

Pollination
Wind and insects are both important pollination mechanisms for Arceuthobium. There
are no specialized pollinators of dwarf mistletoe. Instead they rely on general
pollinators and wind. Hundreds of insect species have been identified as carriers of dwarf
mistletoe pollen, but only a few dozen have been identified as being important pollinators
26, 50. These pollinators include dipterans (e.g. flies), hymenopterans (e.g.
ants) and coleopterans (beetles). The importance of dwarf mistletoe flower parts (e.g. the
pollen and nectaries) as a food source for insects is not well studied. For those seeking
more information, several excellent reviews on the pollination biology of dwarf mistletoe
have been published 66, 25, 67.

Figure 3. Generalized six year life cycle of dwarf
mistletoe. Modified from Hawksworth and Wiens (1972).

Tree host selection
Whether some trees are more susceptible than others to dwarf mistletoe infestation has
long been a subject of interest to silviculturalists. The dwarf mistletoe of the Southwest
are fairly host specific, that is, one type of mistletoe only has one primary host.
However, there are secondary hosts. For instance, A. americanum will sometimes
infect ponderosa pine even though its primary host is lodgepole pine. Likewise, spruce and
true firs will occasionally host A. douglasii even though Douglas fir is the
primary host. A. vaginatum, which primarily infects ponderosa pine, occasionally
infects lodgepole pine 31.

Large areas of apparently susceptible primary hosts are immune to mistletoe infection.
For instance, ponderosa pines in the Black Hills area of south Dakota are uninfected by
A. vaginatum, while A. douglasii is generally absent from the Douglas fir
forests of the West Coast. Why these areas are immune to mistletoe infections is unknown,
but it is thought to be related to climate 61.

Climate is also thought to influence the distribution of dwarf mistletoe on a more
local scale. Studies suggest that dwarf mistletoe infestations in ponderosa pine forests
are most severe on the driest sites, where tree growth rates are slowest , and least
severe in wetter habitats 18, 43. Also, A. americanum is absent form the
higher ranges of lodgepole pine, apparently because of low temperatures, while A.
vaginatum is absent from lower elevation ponderosa pine forests, presumably because of
high summer temperatures 75. Topography also affects the distribution of dwarf
mistletoe, as they are most abundant on ridges, upper slopes and moderate slopes, and
least abundant on steep slopes and valley floors 43. There is also evidence to
suggest that dwarf mistletoe preferentially infect trees on slopes with west and southwest
exposure, and prefer stands with moderate site indices and low basal areas 43.
Collectively, these studies suggest that dwarf mistletoe preferentially infect stressed or
slow growing trees.

The Effects of Land-Use Practices on the Ecology of Dwarf Mistletoe

Fire suppression, livestock grazing and logging are the primary land use practices that
have altered the ecology of mistletoe in the Southwest. Because these land use practices
have so radically altered the overall condition of Southwest forests, the changed ecology
of dwarf mistletoe is largely reflective of overall changes in Southwest forest
conditions. Fire suppression and livestock grazing have both altered the effect of dwarf
mistletoe by increasing its infection rate, primarily by increasing stand densities and
thus facilitating the direct tree-to-tree transfer of mistletoe. Livestock grazing
increases stand densities by removing grasses and exposing bare mineral soil, which is
ideal for pine seedling germination . Additionally, the hooves of livestock press pine
seeds into the soil, further facilitating germination. Livestock also trample seedlings,
but overall, they promote seedling establishment.

Fire suppression has eliminated the fires that regularly swept through forest
understories 14. Historically, these fires were of sufficient frequency to kill
most pine seedlings, helping to keep recruitment rates and stand densities low. Though
fire suppression is commonly identified as the primary cause of high stand densities,
scientific studies have determined that intensive livestock grazing preceeded active fire
suppression policies by several decades. As a result of this grazing, fire occurance was
greatly reduced long before active fire suppression occured. Other comparative studies
also strongly suggests that livestock overgrazing, not fire suppression, is the primary
factor that has allowed stands to become overstocked 59, 42, 79. Studies of
paired sites where fire has been suppressed in both sites, but where grazing has occurred
in only one site, show that the ungrazed sites have little in the way of conifer
regeneration though fire has been suppressed for many decades (e.g. 125 years). Grasses in
these systems are known to contain allelochemicals (toxins) that can kill ponderosa pine
seedlings 35, 54, further suggesting that competition with grasses is the
primary reason why historically, ponderosa pine recruitment rates were low. Such findings
strongly suggest that land use practices which destroy the native grass understory may be
the primary cause for the rapid increase in pine seedling recruitment. Fire suppression
may have played a secondary role in increasing seedling recruitment, as historically,
fires only eliminated those seedlings that managed to grow where competition with grasses
was minimal.

Past and current logging practices have also contributed to the spread of dwarf
mistletoe in Southwest forests. Disturbing soils during timber harvesting, encouraging
afforestation at tree densities well above historic levels, and selective harvesting have
all contributed to a decline in forest health and encouraged the spread of mistletoe. When
soils are disturbed and ground cover is removed during logging operations, stand densities
increase because pine seedlings readily germinate and grow on bare mineral soil. Such
disturbances are particularly prevalent when heavy machinery is used extensively during
timber operations. In general, silviculturalists do not consider such disturbances a
problem because undisturbed forests of the Southwest are considered to be stocked at
numbers far below their productive capabilities. Thus any activity that artificially
increases stand densities, whether it be fire suppression, livestock grazing, or logging
activities, is considered beneficial.

Selective tree harvesting that leaves infected overstory trees standing, also
contributes to the spread of mistletoe. These large trees rain seeds down upon the new
stand that is emerging below, thus infecting an entire new generation of trees, and
ensuring that their lives are short. Recognition of this overstory-to-understory
transferal of mistletoe infections have lead foresters to conclude that removal of such
large trees is the best way to prevent dwarf mistletoe from spreading. However, such
thinning practices fail to consider that under normal conditions, there would be minimal
tree regeneration (and thus minimal infections) near infected trees, because of fires and
competition from grasses. It is more accurate to consider the unnaturally high numbers of
regeneration trees as the problem contributing to the spread of mistletoe. Removing the
large trees creates a forest that is quite unlike anything that historically existed. This
thinning technique, where the largest, most ecologically important trees are removed from
the forest, is known as sanitation salvage, and is being practiced by the Forest Service
as way to improve ecosystem health. However, the idea that ecosystem health can be
restored by simplifying forestsremoving the most important large structural elements in a
forest and replacing them with a stand of even aged seedlingsis counter intuitive and
ignores basic ecological principles. It is a basic tenet of ecological theory that
structurally complex habitats are more diverse than simplified habitats 34, and
habitats that are biologically diverse are generally healthy.

If the purpose of removing infected trees is to improve ecosystem health by preventing
the spread of mistletoe, then the infected trees can simply be killed and left standing,
as mistletoe will not survive on dead trees. Or better yet, simply creating a small donut
shaped no tree zone around infected overstory trees, to serve as a barrier to mistletoe
seed dispersal, should solve the problem without further impoverishing the forest.

Logging has not only increased the spread of mistletoe in small trees, but has also
removed the ecologically important brooms created by dwarf mistletoe infections in large
trees. Unfortunately, current forest practices encourage the removal of these structures.
Even when the tree are not cut, thinning and pruning operations designed to save stands
(so they can later be harvested) typically remove infected trees and limbs from infected
trees. These tree and limbs often contain witches brooms, and many of the trees, because
they are infected, will soon be snags. Thus, these well intentioned clean up efforts to
improve the health of trees, actually reduce the structural elements of a forest that most
greatly contribute to overall ecosystem health and biodiversity.

MISTLETOE "CONTROL" AND THE
DISAPPEARANCE OF LARGE TREES

The Elk Timber Sale

The Elk Timber Sale, on the Lakeside District of the Apache-Sitgreaves National Forest
in eastern Arizona, is a classic example of mistletoe control being used as a rationale
for cutting large trees. Because of previous logging, the Lakeside District has no
remaining old growth at all. The few moderately sized trees which do exist are clumped in
pockets of 3-10 trees surrounded by acres of young pines. Though much smaller than the
giant pines which once blanketed this area, the largest trees remaining are still of
exceptional wildlife and recreation value. On the Elk Timber Sale, trees larger than 16
inches dbh make up just 3% of the landscape, while trees less than 9 inches dbh cover 70%
of the area (Fig. 4).

Figure 4. Although large trees make up a small percent of all trees in the
Lakeside District, they continue to be disproportionately harvested at a non-sustainable
rate.

With the Elk Timber Sale, however, the Forest Service chose to log many of the rare
large trees for mistletoe "control" and to encourage the growth of vast numbers
of small trees. According to their Environmental Assessments, "Large ponderosa pines
can not only spread mistletoe downward if infected, but can effect growth of the
understory (i.e. small trees) by shading and utilizing water and nutrients with the
massive root systems." And, "Removal of infected trees will allow residual trees
to live longer, allowing nearby stands with smaller diameter trees to grow faster..."

Tragically, many of the large trees logged for mistletoe control were in old growth
reserves (mature forests to be left undisturbed so they can develop into old-growth
forests) and protected Northern goshawk nesting territories. Twenty seven percent of the
trees did not even have any verifiable infection. The Forest Service claimed that these
trees, had latent infections, or were in excess of the number of large trees needed, even
though the District has no old growth and is deficient in all large tree classes.

Integrated Management Strategies for Controlling Dwarf Mistletoe

Although dwarf mistletoe is frequently referred to as a problem, that needs to be
controlled, it is more correctly viewed as a component of Southwest forests that is
increasing in abundance because of land use changes that have occurred in the past 125
years. The spread of mistletoe is symptomatic of the decline in the health of Southwest
forests that has occurred because of these changes. Specifically, livestock grazing, fire
suppression and certain logging practices have all contributed to the decline in forest
health and to the spread of dwarf mistletoe. An integrated management strategy that
restores some of the fundamental components and processes that historically existed in
these systems would largely eliminate the mistletoe problem. We refer to this strategy as
integrated because the components (outlined below) are interrelated. All components need
to be incorporated into an overall management plan for any one of them to work correctly.
Such an integrated strategy would include the following fundamental components:

No cutting of large diameter trees and snags. There is no justifiable ecological
or forest health reason for removing large trees or snags. Mistletoe infections are a
normal ecological process whereby large trees are converted into snags. The live infected
trees today are the snags of tomorrow. Unfortunately, large trees and snags are relatively
rare components of Southwest forests because of past and current timber harvesting
activities. There really are very few truly large trees left. The smaller ranges of trees
today classified by the Forest Service as large (17 in > dbh < 24 in) 36
were not even considered to be merchantable less than 90 years ago 3. However,
the largest existing trees do provide the best current habitat, and will provide a future
source of large snags. Therefore, they should not be removed. If a tree is infected with
mistletoe and it is determined that the tree should be killed because it poses a high risk
of infection to other ecologically important trees, then it should be killed and left
standing, to serve as snag habitat.

Thin understory trees to create stand structure and densities that approximate
presettlement conditions. The overstocked young forests of today facilitate the spread of
dwarf mistletoe and in some instances, present a fire hazard to large, ecologically
important trees. Any thinning operation, however must leave the largest trees in a stand
intact. The current Forest Service practice of removing large overstory trees during
thinning operations is an ecologically destructive practice that degrades ecosystem
health. Understory thinning will slow tree-to-tree spread of dwarf mistletoe or limit it
to isolated clusters of trees. All downed small diameter trees and slash should be removed
to reduce the forest floor fuel load and minimize fire hazards. Current and historic
logging practices which leave large volumes of slash in the forest increase fire threats
and beetle infestations.

Reestablish regular ground fires in order to minimize seedling survival and to
prevent the accumulation of fuel. Although crown fires did occur on occasion, the
historical record suggests that they were rare in ponderosa pine forests. Hence, crown
fires should not be encouraged. Before fires are reintroduced, forest understories need to
be thinned and accumulations of forest floor fuels must be removed to prevent intense
fires from killing the few remaining large trees. Although fires, if controlled properly,
will improve forest health, it is essential that large trees be protected or the
ecological benefits that occur because of burning may be outweighed by the loss of the big
trees that take centuries to replace. Given the rarity of large trees, it is extremely
important to remove fuel near their bases to prevent their untimely demise. The frequency
of ground fires should approximate presettlement fire frequencies.

Reduce livestock densities to a level that will allow a relatively continuous
ground cover of herbs and grasses to develop where light, soil and moisture conditions
would normally support such vegetation. Once forests are thinned and opened up, they will
simply return to their pre-thinning densities if livestock remain to prevent the
reestablishment of ground cover. In areas where the ground cover has been eliminated,
livestock should be completely removed until the vegetation has recovered. Future
introductions of livestock in such areas should occur on a limited basis and under careful
monitoring. Should the ground cover begin to degrade, livestock should be removed again
until vegetative recovery is complete. In general, livestock should only be grazed in
forests where there is a relatively complete ground cover, and in these areas, only at
densities and frequencies low enough to prevent increases in the extent of exposed mineral
soil. Experimental evidence from other arid systems (e.g. eastern Oregon) suggests that
removing livestock from degraded areas will improve foraging opportunities 22.
This suggests that removing livestock from damaged Southwest forests may, in the long run,
improve ecosystem health and provide more livestock forage.

MISTLETOE "CONTROL" AND THE DISAPPEARANCE OF
LARGE TREES

The Rocker Timber Sale

In 1991, the Forest Service announced plans to log a portion of the Gila National
Forest in New Mexico, in what became known as the Rocker Timber Sale. Dwarf mistletoe, the
announcement stated, was not a severe problem. A year later, the Forest Service
unexpectedly announced that mistletoe was a severe problem on over 1,600 acres. Overnight,
the Rocker Timber Sale was transformed into a mistletoe "control" project, and
shortly thereafter the largest trees were logged under the guise of disease control.
According to the Forest Service, approximately 90% of the area had previously been logged,
virtually all the old growth was gone, there were very few large snags left and this
habitat type was "on the threshold of losing community viability."

Prior to the sale, Forest Service biologists suggested that, "A long-term
commitment will very much need to be made to work toward restoring this habitat
community." They went on to predict the ecological impacts of logging in this area,
stating that, The most likely nest trees would be those older-aged residual overstory
ponderosa pine trees and the more disfigured older-aged dwarf mistletoe infected pine and
mixed conifer trees. These trees are the most likely future snags and would be heavily
removed..."

and that, "The potential for snag recruitment should decrease substantially.
Stands remaining in the 4.3 wildlife habitat capability class are projected to decrease by
an estimated 91% in ponderosa pine stands and 47% in mixed conifer stands."

Forest Service managers ignored the advice of biologists, and instead chose to maximize
short-term timber production, admitting that mistletoe control was used to justify this
goal. Six and half million board feet were logged on 2,564 acres. More than 12,650 trees
over 16 inches dbh were cut. Rather than improving the health of the forest, this
mistletoe management project further impoverished an area that had already been severely
damaged by past logging activities.

Summary

The spread of dwarf mistletoe throughout Southwest forests is linked to the overall
decline in forest health that has resulted from 125 years of excessive grazing, logging
and fire suppression. Most problems associated with dwarf mistletoe can be solved by
managing these forests for ecosystem integrity rather than timber protection or livestock
forage. Much of the spread of dwarf mistletoe is directly attributable to artificial
attempts at increasing timber yields above historic levels. Fire suppression, livestock
grazing and logging have all contributed to the spread of dwarf mistletoe. Although there
are legitimate concerns about the spread of dwarf mistletoe, management plans must
recognize the importance of these plants to forest ecology. Dwarf mistletoe is an integral
component of Southwest forests, providing food, shelter and nesting sites for wildlife,
and in general increasing biodiversity. Land managers need to focus on maximizing the
ecological benefits of dwarf mistletoe, while researchers need to learn more about the
ecology of these species. Current attempts to improve ecosystem health by removing
mistletoe-infected overstory trees does more harm than good, and the practice should be
discontinued. An integrated ecosystem management strategy that restores the natural
processes and stand conditions that historically occurred in these forest is the best
approach for controlling dwarf mistletoe while simultaneously restoring ecosystem health.